Pakistan J. Zool., vol. 48(3), pp. 907-911, 2016.

Short Communication

Influence of Matrix Habitats on the Occurrence ofTerrestrial Mammals in Planted Forest Landscapes
Kosuke Maeda,1 Shinsuke Koike,2,* Mina Murao,1 Takuya Ishigane,1 Shota Harasawa,3Takashi Masaki,4 Masashi Soga,5 Tomoko Naganuma2 and Akira Sato1
1Tokyo University of Agriculture, 1-1-1 Sakuragaoka, Setagaya, Tokyo 156-8502, Japan
2Tokyo University of Agriculture and Technology, 3-5-8 Saiwai, Fuchu, Tokyo 183-8509, Japan
3
Kyoto University, Kitashirakawa Oiwake-cho, Sakyo-ku, Kyoto 606-8502, Japan
4Forestry and Forest Products Research Institute, 1 Matsunosato, Tsukuba, Ibaraki, 305-8687, Japan
5Hokkaido University, Nishi 9, Kita 9, Kita-ku, Sapporo 080-8589, JapanAB STR ACT
We studied the responses of terrestrial mammal community to high levels of habitat fragmentation
caused by forestry plantations in central Japan. Our aim was to understand how terrestrial mammals
select remnant natural forest (broad-leaved forest) patches in a fragmented forestry plantation
landscape. Camera traps monitored remnant 12 broad-leaved forest patches for 7 months. Thesepatches differed in the ratio of broad-leaved forest to the surrounding forestry plantation matrix area.
In total, 144 photographs captured. A positive relationship was found between the number ofphotographs of mammals and the area of broad-leaved forest around the remnant broad-leaved forest.
Hierarchical variation partitioning also showed that BF had a substantially greater independent
explanatory power than CF. Future studies of landscapes resulting from different matrix types areneeded to help land managers understand the influences of habitat configuration on patterns ofspecies persistence and community dynamics.

Forest loss and fragmentation are two importantinteracting processes that may negatively affectbiodiversity (Fahrig, 2003). They both isolate remnant
patches, increasing the risk of extinction for somemammal species (e.g., Bodmer et al., 1997) and reducing
mammal species richness (e.g., Cullen et al., 2000).
Recent studies on the effects of forest fragmentation
showed that fragmented landscapes are complex andvariable systems, because the quality of remnants or theaffinity to the matrix, the dominant component in thelandscape, often strongly influences their functionalconnectivity (e.g., Laurance and Bierregaard, 1997;
Bierregaard et al., 2001). Additionally, several studies
have described how the surrounding matrix influences
populations in forest fragments (e.g. Hinsley et al., 1995;
Stouffer and Bierregaard, 1995).
The matrix can be important because it bothinfluences mammal movement through the landscape(Revilla et al., 2004; Bender and Fahrig, 2005) andprovides potential food resources (Tubelis et al., 2007;
Harper et al., 2008). Thus, knowledge of the effects ofanthropogenic disturbances and matrix quality andstructure on ecological processes are crucial to_______________________________*Corresponding author: koikes@cc.tuat.ac.jp0030-9923/2016/0003-0907 $ 8.00/0
Copyright 2016 Zoological Society of Pakistan

Article Information
Received 6 May 2015
Revised 2 November 2015
Accepted 12 December 2015
Available online 1 May 2016
Authors’ Contributions
SK and TM conceived the idea. KM,
SK and TM contributed to the study
design. KM, SK. MM, TI and SH
collected the data. MS and TN
analyzed the data and wrote themanuscript with input from allauthors.
Key words
Camera trap, forest fragmentation,
forestry plantation, temperate forest,
terrestrial mammals.

understanding patterns of mammal species distribution
and community dynamics in fragmented landscapes(Santos-Filho et al., 2012).
The area covered by planted forest landscapes,
forestry plantations, is one of the common landscapes inseveral countries such as Japan and Australia (Forest
resource assessment, 2010). Therefore, understanding
how to conserve native biodiversity in the remnant
natural forest patches within planted forest landscapes is
an increasingly important issue. The common matrix
environments under this land-use type may provide new
food sources and some shelter for some animal species
that do not use mature natural forest, allowing them tooccupy the matrix or disperse through it (Anderson et al.,2007). Thus, management of the planted forest landscapematrix for biodiversity is easier than that of other matrix
types such as urban or agricultural landscapes.
Nevertheless, the responses of highly mobile mediumsized and large-bodied vertebrates to habitat disturbanceand matrix quality are not completely understood.
In this study, our objective was to understand how
terrestrial mammals select remnant natural forest (broadleaved forest) patches in a fragmented forestry plantation
landscape in Japan. We used the camera traps because it
was useful noninvasive monitoring tool (e.g., Wang et al.,2014). We investigated the effect of matrix habitat
structure on mammalian communities in a hyper-mosaic

908

K. MAEDA ET AL.

landscape between broad-leaved forests and conifer
forestry plantations of Japanese cedar (Cryptomeria
japonica), Japanese cypress (Chamaecyparis obtusa), andJapanese larch (Larix kaempferi) in central Japan.
Methods
Study area
This study lies within the southern part of theAbukuma Mountains, Honshu, Ibaraki Prefecture, central
Japan (36°56′N, 140°35′E, 610–660 m a.s.l.). The meanannual precipitation and temperature are about 1,910 mm
and 10.7°C, respectively, with mean monthly temperature
ranging from −0.9°C in January to 22.6°C in August,based on data from the Ogawa meteorological station(36°54′N, 140°35′E) (Moriguchi et al., 2002). Themaximum winter snow depth is about 50 cm.
In Japan, forests cover 67% of the land area,
including 40% covered by forestry plantations (ForestAgency, 2012). Most forestry plantations and naturalforests form a mosaic and complex structure within thelandscapes of Japan. Conifer forestry plantations cover
more than 60% of forest area in Ibaraki Prefecture (ForestAgency 2012). In our study area, forestry plantations wasdominant vegetation type, however natural forests wereremain partly. Thus, conifer plantations and naturalforests of broad-leaved trees form a complex mosaic
landscape. Fagus japonica, Castanea crenata, Quercus
serrata, Carpinus laxiflora, and Prunus verecunda
dominate the natural forests. To investigate effect of theamount of natural forests among the forestry plantation
matrix on mammal fauna, we measured the amount offorest (forest area) surrounding the remnant broad-leaved
forest at radii of 126, 178, 252, 309, 357, and 399m fromthe mapped points, which correspond to 5, 10, 20, 30, 40,
and 50ha, respectively. We selected 12 natural forest
patches in our 10 km × 15 km study area depend on thematrix condition. Each patch was separated from at least
1.3km (Fig. 1). Among the matrix (surrounding area ofthe selecting 12 patches), forestry plantations weredominant vegetation type, however natural forests werealso part of the matrix. These selected patches differed in
the ratio of natural forest to the surrounding matrix area
(Supplementary Table I).
Camera trapping
The presence of mammals in 12 forest patches wasassessed from May to November, 2012 using infra-red
triggered cameras (5.0 Megapixel Infrared Digital
Motion-detection Camera, National Geographic Store,Margate, FL, USA) in each forest patch. Because thespecies differed in birthing, pregnancy, and dispersalseasons, we monitored mammals for several months
using a total of 214 camera days per study site (1 camera
× 214 d). Each photograph was stamped with the timeFig. 1. Location of 12 forest patches in
Ibaraki Prefecture, Japan, indicated by numbers.
Black, gray, and white denote broad-leaved forest,
conifer plantations, and other land uses, such as
farmland or pasture, respectively (Modified from
Ministry of the Environment Japan, 1999).

and date. The time delay between photographs was set toa minimum of 1 minute. However, we decided thatphotographs less than 1 hour apart would not beindependent. Only independent data should be enteredinto the analysis. At each site, the camera trap was tied toa tree 50 cm above the ground to take a picture ofterrestrial medium-large size mammals, we have nottargeted small mammals such as rodents. At each patch,
we placed the camera traps near the center of each patch
calculating by GIS where the probability of mammal
detection was high (e.g., animal trails) but we did not use
baits. The vegetation of each camera setting wasrepresentative type of each forest patch. We checked eachcamera and changed the batteries monthly. Photo results
were logged into a computer with frame number, date,time, and contents. Mammal species were identified bybody size and body color.

INFLUENCE OF MATRIX HABITATS ON OCCURRENCE OF FOREST MAMMALS

Table I.-Results of camera trapping.

Plot No
123456789101112909Badger

BoarHare

01101
10000000126
1116
7
10000140010000001010

Number of photographs
FoxRaccoon dog

Statistical analysis
In this study, there was a significant correlation
between broad-leaved forest area surrounding matrix area(BF) and conifer plantation area surrounding matrix area
(CF) (5ha: r = -0.84; 10ha: r = -0.90; 20ha: r = -0.93;30ha: r = -0.93; 40ha: r = -0.92; 50ha: r = -0.90, for all P< 0.001, see Table I). We therefore used a hierarchicalvariance partitioning analysis in order to computeindependent contributions of BF and CF accounting forcorrelation between these variables. In doing so, we usedthe ‘hier.part package’ version 1.0–4 in R statisticalsoftware (ver. 3.2.1, R Core Team, 2015). Hierarchicalvariation partitioning was conducted for each spatial
scale (5, 10, 20, 30, 40, 50ha). In the analysis, we analyze
mammal community as a whole rather than by species
because few animals were recorded.
Results
In total, 144 independent photographs were taken ofseven species: wild boar (Sus scrofa), Japanese badger(Meles meles), red fox (Vulpes vulpes), raccoon dog(Nyctereutes procyonoides), Japanese marten (Martesmelampus), Japanese hare (Lepus brachyurus), andJapanese squirrel (Sciurus lis) (Table I). All camera-traps
functioned fully throughout the whole study period.
Individuals could not be recognized for all seven species.
A positive relationship was found between thenumber of photographs and the area of broad-leaved
forest around the remnant (only area within 50ha)(Spearman's correlation, area within 5 ha: r = 0.40, P=0.2,
area within 10 ha: r = 0.54, P=0.07, area within 20 ha: r =0.43, P=0.3, area within 30 ha: r = 0.45, P=0.14, area
within 40 ha: r = 0.48, P=0.15, area within 50ha: r = 0.59,
P < 0.05). Hierarchical variation partitioning showed that
BF had a substantially greater independent explanatory
power than CF for each spatial scale (Table II).

020000000000

Table II.-

MartenSquirrel

0100100000000410300000400
50021000000The independent contribution (given as
the percentage of the total explainedvariance) of BF and CF from hierarchicalvariation partitioning.

Variables50ha
BF: Area of broad-leaved forest
CF: Area of conifer plantation forest
40ha
BF: Area of broad-leaved forest
CF: Area of conifer plantation forest% independent
contribution

95.6
4.4
67.7
32.3

30ha
BF: Area of broad-leaved forest
CF: Area of conifer plantation forest

93.5
6.520haBF: Area of broad-leaved forest
CF: Area of conifer plantation forest

62.4
37.610ha
BF: Area of broad-leaved forest
CF: Area of conifer plantation forest

95.2
4.85ha
BF: Area of broad-leaved forest
CF: Area of conifer plantation forest

99.1
0.9Discussion
Human land uses that displace and fragment forest
habitats serve as important factors in the decline offorest-dependent fauna (e.g., McAlpine and Eyre, 2002;
McGarigal and McComb, 1995; Rochelle et al., 1999).
However, fragmentation by forestry plantations does notnecessarily decrease the total forest area in a landscape.
However, our results indicate forestry plantations aroundthe remnant broad-leaved forest cannot be a substitute for910

K. MAEDA ET AL.

remnant natural forest.
In our study, a positive relationship was foundbetween the number of photographs, especially wild boar,
and the area of broad-leaved forest around the remnant
broad-leaved forest, especially more large scale. And,
there may be ecological threshold values of natural forest
area around the remnant natural forest within a forestry
plantation matrix for the mammal abundance. For
example, very few mammals were recorded when naturalforest is less than 22 ha in the case of the area of broadleaved forest around the remnant natural forest within 50
ha (Table I). However, there are several exceptive patchessuch as No.1 or No. 11. Several plausible reasons may
explain these phenomenons. Firstly, major photographed
species are mid- to large-sized mammals, these species
have large home ranges, which are 116 ha for wild boar,
5–407 ha for badger, and 10–600 ha for raccoon dog(Ohdach et al., 2009). Thus, because these three species
are highly mobile, they may move through much broader
areas. Secondary, the matrix may act as some functions
such as food resource. For example, in particular, thewild boar mainly eat earthworms (Ohdach et al., 2009).
However, the amounts of earthworms do not differ
between broad-leaved forests and conifer plantation
forests (Ichikawa et al., 2008). Third, our study design
has some problems; camera traps do not provide large
amount of data (low number of camera sites and norepeat measures over a number of study years) andindividual recognition is impossible. Thus, it is possiblethat some individuals (especially the larger species) could
have moved/disperse between patches (especially ones
close together). For the future, we plan to investigate howthese animals move between fragments using telemetry.
However, we can provide several importantsuggestions. Previous studies identified matrix quality as
an important factor that determines the persistence ofspecies in fragmented habitats (Bentley et al., 2000;
Knight and Fox, 2000), it may also affect habitat quality
via edge effects, although exotic species and alteredclimatic conditions at patch boundaries affect habitats(Saunders et al., 1991). However, in this study, theconifer forestry matrix may provide additional foodsources and some shelter for mid- and large sized
mammals. We cannot discuss this concept in detail. Thus,
studies of landscapes resulting from different matrix
vegetation structure are needed to aid our understandingof the effects of habitat configuration on patterns ofspecies persistence and community dynamics in thefuture.
Acknowledgements
We thank Mr. Yamagata for field assistance. We
also thank the Ibaraki District Forestry Office for use of

their facilities. The G2P08 research program of theForestry and Forest Products Research Institute and theJapan Society for the Promotion of Science Grant-in-Aidfor Young Scientists (B) 25850103 and Grant-in-Aid for
Scientific Research (A) 25241026 supported this study.
Statement of conflict of interest
Authors have no conflict of interest.
Supplementary Table I is available on Weblink:
http://www.zsp.com.pk/pdf48/907911%20(41)%20QPJZ-219-2015%2019-4-16_skR.pdf
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